Rare or underestimated? The distribution and abundance of the pond bat (Myotis dasycneme) in Poland

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1 Rare or underestimated? The distribution and abundance of the pond bat (Myotis dasycneme) in Poland Mateusz Ciechanowski 1, Konrad Sachanowicz 2 & Tomasz Kokurewicz 3 1 Department of Vertebrate Ecology and Zoology, University of Gdańsk, Legionów 9, Gdańsk, Poland, matciech@kki.net.pl 2 Department of Animal Ecology, Nicolaus Copernicus University, Gagarina 9, Toruń, Poland 3 Department of Zoology and Ecology, Wrocław University of Environmental and Life Sciences, Kożuchowska 5b, Wrocław, Poland Abstract: The paper summarises the distribution and abundance of the globally vulnerable and nationally endangered pond bat (Myotis dasycneme) in Poland and is based on all the available published and unpublished data. The species was recorded in 105 UTM squares, scattered across almost the whole of Poland. In addition fossil remains, mostly from the Holocene age, were found in 13 squares. Evidence of pond bat breeding (nursing) was obtained from 10 UTM squares, restricted to the lakelands of northern Poland and the valleys of the large rivers in the central part of the country. Only two nursery roosts, used respectively by a maximum of 481 and 391 individuals, were found. The remaining summer records (41 UTM squares) were sightings or captures of adult males, non-breeding females or unsexed individuals and were widely dispersed in areas located far from known reproduction sites. The summer roosts (n=21) were located mainly in buildings, bridge crevices and bird and bat boxes. Sixty-seven hibernation sites of pond bat were found, mainly in old fortifications, cellars and caves. 63% (n=42) of them were used by single individuals and only three held 10 or more individuals (maximum n=34). In the two regularly monitored sites the number of pond bats has fluctuated in recent years and in one site it even increased. Most of the available bat data (winter censuses, bird and bat boxes, owl pellets, road casualties, fossil remains, mist netting) shows that pond bats account for less than 1% of the Polish bat assemblages. However this figure reached % (and in one instance 17.5%) of the total number of bats netted in some regions of the northern lakelands. Many areas that are potentially attractive for breeding populations of pond bat have not yet been surveyed, suggesting that the species in Poland may be much more common and numerous than previously thought. Keywords: bats, Chiroptera, Vespertilionidae, endangered species, conservation status, riparian habitats. Introduction In Europe the distribution range of the pond bat (Myotis dasycneme Boie, 1825) extends from southern Sweden, Finland and Russia in the north, to Belgium and France in the west and Croatia, Serbia, Bulgaria and Ukraine in the south (Horáček 1999, Limpens et al. 2000, Benda et al. 2003, Siivonen & Wermundsen 2003). A vagrant individual was recently also found in Great Britain (Hutson 2005). Within this distribution range, 2007 Vereniging voor Zoogdierkunde en Zoogdierbescherming. Lutra articles also on the internet: reproduction (i.e. nursing) has been observed in the Netherlands, Denmark, northern Germany, Hungary, Latvia, Estonia and Russia (Limpens et al. 2000). The pond bat is a stenotopic species that prefers a lowland landscape with a dense network of patches of still or slow-moving freshwater and uses a specialised hunting tactic (trawling). Its continued survival is a high conservation priority (Horáček & Hanák 1989, Limpens et al. 2000) and it was included in the 1996 IUCN Red List of Threatened Animals and classified as vulnerable (VU) in the Palearctic Region (Hutson et al. 2001). As a result the pond bat is included in Appendix II of the EC Directive on the Conservation of Natural Habitats and of Wild Fauna and Flora Ciechanowski / Lutra (2):

2 (92/43/EEC) as a species that demands creation of special areas of conservation (SAC) within the Natura 2000 Network. It is legally protected at the international level through the Bern Convention on the Conservation of European Wildlife and Natural Habitats, the Bonn Convention on the Conservation of Migratory Species of Wild Animals and the Agreement on the Conservation of Populations of European Bats (EUROBATS) and is nationally protected in all European countries. While generally considered as a rare, patchily distributed and seriously threatened species in Western and Central Europe, it is one of the most abundant bat species within its Russian range (Strelkov & Iljin 1990, Bolshakov & Orlov 2000). In 1986 it was thought that the population of the pond bat was as small as 3,000 in Western Europe and that the world population was about 7,000 individuals (Stebbings & Griffith 1986). More recently the number of pond bats in countries designated as population centres, i.e. Hungary, Latvia, the Netherlands and Russia, has been estimated as between ,000 (Limpens et al. 2000), indicating a substantial earlier underestimation. This discrepancy was largely caused by a lack of surveys on the occurrence and abundance of the pond bat over most of its range (Limpens et al. 2000). In Western Europe, the only available data concerning long-term population changes are from the Netherlands. Population trends based on calculations by Statistics Netherlands (CBS) indicate that the numbers in hibernation sites have more than doubled between 1986 and 2005 (Verboom 2006). A similar increase was also found earlier in Estonia (Piusa Caves; Lutsar et al. 2000) and recent information from other parts of Europe within the range of the pond bat indicate that the numbers are stable or slightly increasing (e.g. Baagøe 2001). In Poland the pond bat has traditionally been treated as a rare species confined to lowland areas with an abundance of water (Kowalski 1955, Kowalski & Ruprecht 1981). The first comprehensive review of the distribution of Polish mammals listed 33 localities of pond bat (Ruprecht 1983) but made no clear distinction between winter and summer locations. About half of the distribution data were drawn from pond bat remains found in owl pellets. There were no confirmations of the species breeding in Poland in the 20th century even though it was suspected that this occurred in some areas. An earlier claim by Horáček and Hanák (1989), that Krzanowski (1956) provided direct evidence of the reproduction, appears to be a misunderstanding (Ciechanowski et al. 2002). Horáček and Hanák (1989) also speculated that the pond bat avoids large areas of the Polish lakelands (e.g. the Masurian and Pomeranian regions) as it prefers linear water bodies (canals, blind river branches) rather than large lakes. The Action Plan for the Conservation of the Pond Bat Myotis dasycneme in Europe (Limpens et al. 2000) estimated roughly 1,000 individual pond bats in Poland. The species is classified as an endangered species (EN) in the National Red Data Book (Wołoszyn 2001) but the statement that it had strongly declined in recent years is not supported by any literature or original field data. Recently, localities of pond bat have been identified in almost all regions of Poland (Sachanowicz et al. 2006) although there has been no up-to-date review of the species distribution and status in the country, which is essential for planning national conservation measures. This review seeks to fill this gap by presenting the current state of knowledge on the distribution, abundance and status of the pond bat in Poland. Material and methods In describing the distribution of the pond bat in Poland, we have used all the available data, including both published (81 publications, 262 records) and unpublished reports (39 Polish and foreign bat workers, 88 records). These include both recent and historical information (from the C19th to June 2007). This information was obtained trough a variety of methods: mist netting, surveys of summer and winter roosts, radio-tracking individuals captured in their summer colonies, ultrasound detection and observations of foraging or commuting bats and 108 Ciechanowski / Lutra (2):

3 Table 1. The total number of localities and UTM squares, where the pond bat has been recorded. 1 Nurseries, captures or findings of lactating females and juveniles in May August; 2 Swarming sites were included, however all of them appeared to also be hibernacula. March April and September October were considered as transitional periods. Type of record Number of localities Number of UTM squares summer breeding summer non-breeding winter and transitional quarters undetermined period owl pellets museum specimens 5 5 total collecting dead specimens and owl pellets (see Limpens 2001 for review on methods). Here we apply the terms breeding and reproduction as synonyms for nursing, so breeding is only acknowledged if a nursery roost, lactating female or juvenile individual was recorded in a particular site. Three localities where the pond bat was claimed to be present were rejected as doubtful, either because the method used was insufficient to recognise the species (Sitowski 1948) or the author did not provide any details about the place, date and method (Čmak 1968). In addition we separately review fossil records of the species. A complete synopsis of all these records and their references is given in Annexes 1-5. All localities were attributed to the particular UTM coordinates, using Gnomon 3.3 software (Desmodus, Poland). We also analysed the use of particular types of roosts used by pond bats, as well as the percentage of pond bats within available bat samples netted, counted in roosts, collected as road casualties, extracted from owl pellets and fossil material (only samples where n>100 individuals were included). Results General and summer distribution In total, the pond bat has been recorded as present at 171 sites, located in 105 UTM squares (table 1). These sites were spread across the whole of the country, with no limit of distribution and no clear distribution pattern (figure 1). Concentrations of records (Warsaw area, the Upper Narew and Biebrza Basins) and those regions where any recorded localities are lacking (including some parts of the Northern Lakelands) were associated with different intensity of bat surveys conducted in these areas. However, some distinct distribution patterns can be found when summer and winter records are separately considered. In summer almost all the records were obtained from the low-lying parts of the country, with only three records from localities in the Southern Uplands and one from mountainous areas (the Sudety Mountains) (figure 2). The lack of records from the Carpathians was not due to a low intensity of survey efforts, but rather to the avoidance of higher altitudes by pond bats in summer. We cannot, however, exclude the incidental occurrence of pond bat in the Polish Carpathians outside of its hibernation period, as the species has been discovered in the Slovakian part of the Tatra Mountains and foothills, both at foraging sites and cave entrances (Pjenčák et al. 2003). Evidence of breeding was found in only ten UTM squares (Annex 1), all concentrated in the lakelands (Masurian, Pomeranian) or large river valleys (Vistula, Warta). Recent breeding records ( ) are restricted to the northern and central parts of the country (figure 2). Two currently active nursery roosts, one Ciechanowski / Lutra (2):

4 Figure 1. Distribution of the pond bat in Poland. Grey circles records before 1939, grey squares records from 1939 to 1983; black circles new records since publication of Ruprecht (1983); black squares sites where the presence of pond bat was recorded before 1983 and confirmed later. Complete list of numbered localities and records in Annexes 1-4. in the wooden church in Jeleniewo (Suwałki Lakeland, UTM FF 20) and the other in an old forester s lodge in Lubnia (Pomeranian Lakeland, UTM XV 88) (Wojciechowski et al. 1999, Ciechanowski et al. 2003a) were used by maximum numbers of 481 and 391 individuals, respectively. Both of these roosts are protected as Natura 2000 sites (PLH and PLH respectively). In the 19th century, a third nursery roost was reported in an old church in Trześnia (Sandomierska Forest, southern Poland, Jachno 1868) but no data about its size are available. The remaining breeding records relate to lactating females and juveniles that were mist-netted above rivers (five localities), feeding grounds of radio-tracked lactating females (two localities), a juvenile killed in a road accident (one locality) and a historical record of a female with a juvenile in a tree hole, probably a satellite roost, in Sokolniki near Trześnia. Analysis of the data (figure 2) indicates the presence of at least three other 110 Ciechanowski / Lutra (2):

5 Figure 2. Summer distribution of the pond bat in Poland. Large black circles breeding sites recorded between 1990 and 2007, large grey circle historical breeding site from 19th century, small open squares non-breeding records. Known and suspected nursery colonies: 1 Jeleniewo, 2 - Lubnia and the environs of Wdzydze Lake, 3 Trześnia, 4 Iława Lake Region, 5 Vistula river valley near Warsaw, 6 Upper Warta river valley; recorded colonies of males: 7 Puławy, 8 Golesze, 9 Żłobek. nursery roosts situated in Iława Lake Region, the Vistula river valley (near Warsaw) and the Upper Warta river valley. The largest concentration of breeding records was noted around Wdzydze Lake (Pomeranian Lakeland, UTM XV 88, 89, 98), where, apart from the nursery colony in Lubnia, 14 lactating females and 6 juvenile individuals were netted over the Wda river at four distant localities (Ciechanowski et al. 2006b). The remaining summer records were mostly of single adult males (both netted and found in roosts) and were widely distributed across the low-lying parts of Poland (figure 2). Non-lactating or non-pregnant adult females were recorded in just four localities. Some individuals found in roosts remained unsexed or the original publications did not provide data about their sex and status. In three instances (Puławy, Golesze, Żłobek), small summer colonies of 9-12 males Ciechanowski / Lutra (2):

6 Figure 3. Winter distribution of the pond bat in Poland. Size of circles reflects maximum number of individuals counted during one census. were found (Annex 2). The distribution of nonbreeding individuals seems to be unaffected by the size and density of water systems and they even occur in the dry lowlands and uplands of southern Poland. In ten summer localities the occurrence of pond bat was determined by the use of ultrasound detectors and observations of flying individuals, mostly in the typically-preferred biotope of large water surfaces. Some historical data on foraging individuals over the fish ponds of Lower Silesia (Pax 1925) and the Dunajec River in the Pieniny Mountains (Sitowski 1948) should be regarded as doubtful (e.g. Kowalski 1955) as they were based exclusively on visual observations. Winter distribution and abundance Hibernacula, spring and autumn transitional roosts and underground swarming sites associated with these bats are widely, although unevenly, distributed across the country (figure 3). In most winter roosts (n=42, 63%) single individuals were found and only in 13 hibernacula (19%) were two bats counted simultaneously. Larger winter sites are even rarer: occurrence of 3-5 in- 112 Ciechanowski / Lutra (2):

7 40 November-December January-February number of individuals Biebrza and Narew river basins (northeastern Poland) Year 12 number of individuals Year Fort Carré of the Wis ouj cie Fortress (northern Poland) 25 number of individuals Year "Nietoperek" bat reserve (western Poland) Figure 4. Changes of pond bat numbers in the largest Polish hibernacula. Data from all the regularly monitored sites in the Biebrza and Narew river basins (Osowiec, Piątnica, Drozdowo, Trzcianne, Downary) were pooled together. Ciechanowski / Lutra (2):

8 dividuals was found in seven roosts (10%), 6-9 individuals in three roosts (4%), while only three sites (4%) were used by 10 or more hibernating pond bats (Annex 3). All the main hibernacula of the pond bat are located in lowland river valleys or in lakeland areas (figure 3). The site with the most wintering individuals is in Fort Osowiec II in the Biebrza Valley (up to 34 bats); with the other large winter sites being the underground corridors of the Nietoperek bat reserve (up to 21) and the Wisłoujście Fortress on the Baltic Sea coast (up to eleven). The biggest concentration of hibernacula and hibernating individuals, consisting of 16 different roosts, is around the swampy surrounds of the Biebrza and the Upper Narew River Basins. These accounted for 9.7% (n=165) of all hibernacula surveyed in that region (Kowalski et al., 2003). The Międzyrzecki Fortified Front accounts for relatively large numbers of pond bats, wintering not only in the main underground fortification system (see above) but also in three smaller bunkers, where an additional ten individuals were found in Several individuals were regularly captured in this area during autumn swarming (Łupicki et al. 2001). Several winter sites of the pond bat have also been found in the karstic uplands and mountainous areas in the south of the country, which is rich in caves and abandoned mines (figure 3). In most cases these contained only 1-2 hibernating individuals. Although the species was recorded in 7.9% (n=132) of the caves in the Kraków-Częstochowa Upland and Sudety Mountains these 16 localities are dispersed across an area more than twice the size of the Upper Narew and Biebrza Basins (Nowak & Kozakiewicz 2000, Postawa & Zygmunt 2000, Furmankiewicz & Furmankiewicz 2002, Nowak et al. 2002). Between 1999 and 2007 the number of pond bats at the Nietoperek bat reserve increased. Strong fluctuations in numbers were observed and between 1993 and 2000 at the five sites in the Upper Narew and Biebrza basins between at Fort Carré / Wisłoujście Fortress (figure 4). Significantly more pond bats were counted in November-December than in January-February in the latter area (Wilcoxon test, T=1.00, Z=2.20, P<0.03). The species hibernates in the Carpathians only incidentally and this exclusively in the highest part, the Tatra Mountains. The highest recorded recent European sightings of pond bat are all located here; in the Magurska, Psia and Miętusia Wyżnia caves (at 1465, 1410, and 1393 masl, respectively Nowak et al. 2001). There are no winter records of the species from any other parts of the Carpathians (e.g. the Beskidy Mountains), although numerous surveys have been conducted in the sandstone tectonic caves in that region (Mleczek 2002), or amongst the winter bat colonies of Western Pomerania (Dzięgielewska 2002, Bernard & Samoląg 2002, Gawlak et al. 2002). Fossil occurrence The distribution of fossil and sub-fossil records (28 localities, 13 UTM squares) is restricted to upland and mountainous regions where cave deposits occur (figure 5). Remains that resemble the pond bat were described as M. dasycneme subtilis from the basal Pliocene of the Kraków- Częstochowa Upland by Kowalski (1956), although this was later considered as conspecific with Myotis delicatus Heller, 1936 (Horáček and Hanák 1989). Pond bat was also found in late Pliocene deposits in two localities in the same region (Annex 5). Remains of the pond bat, which were identical with recent species, were also found in 4 Pleistocene and 21 post-glacial (Holocene) sites in the Kraków-Częstochowa and Roztocze Uplands and in the Świętokrzyskie, Sudety, Tatra and Pieniny Mountains (figure 5). In the Holocene period, pond bats reached even higher altitudes than they nowadays, visiting the caves that are now located above the tree-line in the Tatra Mountains (in the Nad Dachem, Studnia w Kazalnicy and Ptasia Caves at 1570, 1545 and 1627 masl respectively Piksa & Wołoszyn 2001). Recently three Polish caves containing post-glacial remains of the pond bat have been used by the species for hibernation (Postawa 2004). 114 Ciechanowski / Lutra (2):

9 Figure 5. Distribution of fossil remains of the pond bat in Poland. Black circles Holocene sites, grey circles Pleistocene, grey squares sites from the Pleistocene and Pliocene periods, black squares sites from the Holocene, Pleistocene and Pliocene periods. Complete list of numbered localities and records in Annex 5. Percentage of the species in bat assemblages The pond bat appears to be one of the rarest bat species in Poland. In bat assemblages it usually constitutes less than 1% of all bats, irrespective of the sampling method employed. In national banding programmes it accounted for 0.44% of the total bat number in (Kowalski et al. 1957), 0.38% in (Krzanowski 1960) and 0.59% in (Harmata 1996). It constitutes 0.73% of the 1,240 bat remains found in barn owl (Tyto alba) (Ruprecht 1979) pellets and 0.58% of the 172 bat remains in the food of tawny owl (Strix aluco) (Ruprecht 1979, Kowalski & Lesiński 1990, Ruprecht 1990, Kowalski & Lesiński 2002), collected across all of Poland. Among the 4,334 bats found in Polish bird and bat boxes 0.30% were identified as pond bats. Pond bats used this type of roost only in the north-eastern part of the country, where they accounted for 0.93% of 1,401 records (Kowalski & Lesiński 1994). Among the 157 bat road casualties on Polish roads, 1.3% belonged to the pond bat (Lesiński 2007). The species was also rare in bat assemblages in the Holocene period. It accounts for just 0.19% of sub-fossil bat remains found in cave sediments in the Kraków-Częstochowa Uplands (Postawa 2004), 0.34% in those in the Tatra Mountains (Piksa & Wołoszyn 2001) and 0.56% in the Pieniny Mountains (Alexandrowicz et al. 1985, Ochman & Wołoszyn 2003). Pond bat is also an uncommon species during bat winter censuses, constituting about 0.02% of the 139,959 bat winter records between 1988 and 1992 (Wołoszyn 1994) and 0.2% of 1,221 bats hibernating in small village cellars throughout the country (Lesiński et al. 2004). The occurrence of pond bat in winter assemblages during February censuses varied between regions, from % in the caves of the Tatra Mountains (Piksa & Nowak 2000) and the fortifications of the Masurian Lakeland (Fuszara et al. 2002) to 0.15% in caves in the Sudety Mountains (Furmankiewicz & Furmankiewicz 2002), 0.29% in caves of the Wieluń Uplands (Kowalski et al. 2002) and 1.70% in anthropogenic underground sites in the Biebrza and Narew valleys (Lesiński & Kowalski 2002). In post-soviet military bunkers in the Podlasie Lowland pond bats constituted just 0.1% of bats counted between December and February, although it reached 1.8% in September-November (Sachanowicz 2007). Exceptionally high proportions of pond bat were observed in winter samples from the bunkers at the Międzyrzecki Fortified Front, away from the main underground system (2.78%; Szkudlarek et al. 2001), in the November-December sample from Ciechanowski / Lutra (2):

10 the Biebrza and Narew Valleys (5.4%; Lesiński & Kowalski 2002) and in autumn and winter samples from the Wisłoujście Fortress (5.2% and 3.0%, respectively; Ciechanowski et al. 2006a). In the largest Polish hibernaculum, the underground system of the Nietoperek bat reserve, only 0.05% (16 out of 32,200) of bats counted in the 2004/2005 winter season were identified as the pond bat (authors unpublished data). The percentage of pond bats among bats captured in mist nests over water bodies in Poland is generally low. It varies from none in most areas of southern and central Poland (Kowalski et al. 1996, Rachwald et al. 2001, Mysłajek 2002, Mysłajek et al. 2002, Sachanowicz & Krasnodębski 2003) to 0.5% in Western Pomerania (Wojtaszyn 2002) and % in the eastern part of the Northern Lakeland belt (Ciechanowski et al. 2002, Postawa & Gas 2003). The summer assemblage of bats in the Wdzydze Landscape Park (Eastern Pomerania) appears to be unique in Poland, as the pond bat constitutes 17.5% of all bats captured there in mist nets above the rivers and is the second most numerous species recorded by this method (Ciechanowski et al. 2006b). There is little data about the numbers of pond bat among bats swarming at the entrances of underground roosts, figures vary from none in mountainous areas (Furmankiewicz & Górniak 2002, Węgiel et al. 2004) to 1.1%- 3.3% in the Międzyrzecki Fortified Front on the western lowlands (Łupicki & Kowalcze-Łupicka 1999, Łupicki et al. 2001). Habitat use No systematic survey on habitat selection by pond bats in Poland has been conducted, but some conclusions may be drawn from the material reviewed in this paper (Annexes 1-4 in supplementary material). Summer roosts, including all nurseries and male colonies, are mostly located in buildings (lofts, spaces inside roofs and wall cavities: 8 sites). Non-breeding individuals were observed in bird and bat boxes, bridge crevices and, exceptionally, in caves (table 2). The only two observations of pond bats roosting in tree holes were made in the 19th century. One of them, made in August 1861 in a hollow hornbeam tree in Kaskada Park in Warszawa, could have been a mating roost (Wałecki 1881). Pond bats in Poland only hibernate in underground shelters, mainly in fortifications and caves, small cellars (in northeastern Poland), sporadically in mines, underground quarries and once in a village well (table 2). Exceptionally, the species has been found in a greenhouse in winter (Krzanowski 1963) and in a sewer pipe on 17th September (7, unknown locality; Krzanowski 1959b). Observations of copulating individuals, hidden in crevices in the internal brick walls of the Wisłoujście Fortress (autumn 2002) were evidence that some hibernacula may also be used as mating roosts (Sachanowicz & Ciechanowski 2005). Observations and captures of foraging or commuting pond bats have been made on large and medium-sized rivers (n=10), lakes (n=9), small rivers and streams (n=7), fish ponds (n=5), canals (n=4), oxbow river branches (n=2), a campsite near a river (n=1), artificial reservoirs (n=1) and an old park (n=1). Initial radio-tracking observations conducted in Jeleniewo (n=5 ad. tagged) revealed that the feeding grounds were located km from the nursery roost, exclusively over large mesotrophic lakes (Kokurewicz & Furmankiewicz, unpublished data). Discussion Although the pond bat is considered as endangered in Poland (Wołoszyn 2001) it seems to be quite widespread in all of the lowland parts of the country and is recorded as breeding in five areas (figure 2). Other large areas are utilised mostly by non-breeding individuals that may live solitarily and disperse much more widely than breeding females. Another water-surface forager, daubenton s bat (Myotis daubentonii (Kuhl, 1817)) reveals local sex segregation in summer, when pregnant and lactating females occupy areas much closer to optimal foraging sites than adult males (Encarnação et al. 2006). the pond bat appears to express a similar pattern on a much wider geographical scale. This 116 Ciechanowski / Lutra (2):

11 discontinuous distribution of pond bat has also been observed in the rest of its European range. The concentrations of maternity roosts are known in only a few European regions (mainly the lowlands of the Netherlands and Belgium, Eastern Jutland in Denmark, the northern German lakelands and the Tisza river basin in Hungary) which are commonly recognised as the reproduction centres, while there are regular non-breeding bat records from other regions (Horáček & Hanák 1989, Limpens et al. 2000, Van de Sijpe et al. 2004). Occurrence of the species in small, partially isolated, areas as is observed in Western and Central Europe seems to be characteristic only in the edge zone of the species distribution range. It is difficult to decide if the widespread occurrence of the pond bat in Latvia (Pētersons & Vintulis 1998) indicates that this might also be another reproduction centre or rather the Latvian population is an extension of a continuous core range, that also covers large areas of Russia, where the pond bat is one of the dominant species in bat assemblages (see Strelkov & Iljin 1990, Bolshakov & Orlov 2000, Chistyakov 2001). Scattered, spatially restricted populations of habitat specialists are generally recognised to be in serious threat of extinction and it has often been suggested in the past that this was the situation faced by the pond bat, especially as there was evidence that these islands of distribution have shrunk seriously in the past (as indicated by sub-fossil materials from some European countries, Horáček & Hanák 1989). However, the pond bat in Poland seems to be much more common and much less threatened than previously stated (Limpens et al. 2000, Wołoszyn 2001). No long-term population trends of the species could be traced in Poland, largely because national bat monitoring is mainly based on winter censuses of the underground roosts (Wołoszyn 1994) in which only a few dozen individual hibernating pond bats are usually counted. Such a small sample is highly sensitive to stochastic phenomena (e.g. accidental deaths of particular individuals) and cannot reflect any more general trends in the population at large. Moreover, estimating bat numbers from winter roosts, where censuses traditionally take place in January and February, can be inappropriate for the pond bat, which reaches much higher numbers in autumn (Lesiński & Kowalski 2002) and later probably hides in inaccessible places. Wintering pond bats observed in Poland usually hibernate in deep and narrow crevices (authors observations) so might often be overlooked in some site censuses. The total number of individuals inhabiting the two known maternity roosts is several times higher than in all recently visited hibernacula so we do not know where most of the Polish pond bats spend the winter. This situation is very similar to that observed in the Netherlands, where the known summer population of pond bats is 20 times higher than the winter one (Limpens et al. 2000). Overall, there is no evidence to suggest any recent decline of the species in Poland, as the winter counts reveal only fluctuations or even possibly a periodic increase in the numbers of pond bats. There are however, much more important reasons for rejecting the hypothesis about any decline in this species. Firstly, the abundance of the pond bat in Poland seems to be strongly underestimated, as it was previously in the Netherlands, where systematic and goal-directed surveys between 1986 and 1993 revealed a population 4-5 times larger than previously estimated (Limpens et al. 2000). Low dominance indices (<1%) for pond bats obtained from most Polish samples might imply that it is a rare species. Most of the intensive bat surveys, however, have been carried out in regions that do not have the optimal habitat conditions for pond bats. In some localities, the species is both common and numerous, as netting surveys in the Wdzydze area and Biebrza basin have recently shown. Although mist netting appears to be an effective method for pond bat surveys, it may only work well under specific conditions, where medium-sized rivers and canals serve as commuting corridors (cf. Limpens 2001). In other regions that are potentially suitable for the pond bat, surveys need to be based on inspections of lofts and attics and on ultrasound detection, combined with visual observations of foraging individuals. These surveys need to be conducted by people familiar with the species (Limpens 2001). No such studies have yet been carried out in most of the lakeland or Ciechanowski / Lutra (2):

12 fishpond regions of Poland and there has been no systematic bat survey done in either Żuławy Wiślane (delta of the Vistula river) or the delta of Odra river regions, where the landscapes, rich in canals, ditches and blind river branches are suitable for pond bats. In addition, the range of optimal hunting habitats for pond bats is much wider than has been previously suggested (Horáček & Hanák, 1989). Both the known Polish nurseries for the pond bat are located in young, post-glacial lakeland landscapes, which are only rich in lakes (mesotrophic rather than eutrophic) but without any broad canals or slowly flowing, large rivers. Such landscapes, formerly considered as unfavorable for the species, resemble the habitat utilised by the Latvian population (Pētersons & Vintulis 1998) which has one of the highest population densities in the European Union (Limpens et al. 2000). In the light of this knowledge, breeding populations of pond bat can sustain themselves in any areas rich in large water bodies, and it is very likely that other maternity roosts will be discovered in geographically similar other regions of Poland (e.g. the Lubuskie Lakeland, the southern part of Pomerania or Polesie) that have not yet been properly surveyed. Given this possibility and recent records, it is possible that Poland could be recognised as another European reproduction centre in Europe, joining those already identified by Limpens et al. (2000). Equally it may be possible to treat Polish breeding colonies as stepping stones within a larger, continuous, reproduction area that ranges from northern Germany, across the Baltic States and onto the Russian populations. There is however one gap in this distribution, in Lithuania, where despite frequent summer records the only known nursery is in the north-east of the country and possibly represents an extension of the large Latvian population (Mickevičiene et al. 1999). In southern Lithuania, bordering Poland, the pond bat appears to be a rare bat, and there are only a few records of non-breeding bats (Pauža & Paužiene 1999), although these do include the largest hibernaculum in the country, the Paneriai tunnel in Vilnius, used by about 85 individuals (Baranauskas 2006). Any hypothesis about numerous, but locally distributed and mostly undiscovered, summer populations of pond bats in Poland is not necessarily shaken by a lack of large winter aggregations, even though mass hibernation of the species ( in one site) is known from Denmark (Baagøe 2001) and Russia (Bolshakov & Orlov 2000). Only in Danish Jutland, there is evidence of one population of pond bats reproducing and hibernating in the same area (Baagøe 2001). In general the species has strong migratory tendencies (with female movements up to 300 km). A large part of the lowland Dutch population migrates to the hilly area of Limburg and German mountains, where there are optimal hibernacula (Roer 2001). Similarly, Polish summer populations could leave the country for winter. Equally, the Polish main hibernacula could be simply still remain undiscovered because of their observed tendency to aggregate in large numbers in a just few of many available sites (cf. Bolshakov & Orlov 2000). It is not possible to predict whether any natural or anthropogenic factors could lead to a serious decline of the pond bat in Poland in the near future. The pond bat, as a synanthropic species, makes great use of buildings as maternity roosts in the summertime. Recent renovation, rebuilding and demolition activities could be a threat for the pond bat, as old houses, with deep crevices, slowly disappear from the Polish landscape. A second threat could be increasing and uncontrolled disturbance of hibernation sites by tourists. Despite the legal protection that bats are afforded in Poland and the designation of the largest hibernacula as Natura 2000 sites, sight-seeing in underground sites in winter is still possible and in some places, such as the Nietoperek bat reserve, it is steadily increasing (authors unpublished data). Wołoszyn (2001) suggested water pollution as the main threat for the species and this was partially confirmed by field studies by Van de Sijpe et al. (2004), who recorded higher pond bat activity over waterways with mild and moderate pollution than over heavily polluted ones. Yet this result may also be influenced by other un- 118 Ciechanowski / Lutra (2):

13 studied factors (e.g. proximity of roosts) and the methodological issues associated with calculating hunting activity solely from qualitative data (presence/absence in the number of surveys). In polluted areas there is evidence of heavy metals, PCBs and pesticides being absorbed by the aquatic chironomid flies which are the main prey of the pond bat. Dead pond bats, collected in less contaminated areas, contained PCBs at concentrations known to inhibit reproduction among some other mammals (Reinhold et al. 1999). However, water quality has improved in some Polish waters in recent years including in Wdzydze Lake, which is part of the Pomeranian refuge of the pond bat (Żmudziński 1997). On the other hand many of waters in northern Poland are still heavily polluted by untreated sewage effluents. Even if the picture of the status of pond bat in Poland painted here appears more optimistic than earlier thought, this should not diminish conservation efforts. In future, the emphasis of these might change towards focusing on protecting an internationally important population rather than a local peculiarity. The conservation of unevenly distributed species on a European scale should focus on those regions with the strongest and most viable populations, where the pond bat is still common and widespread. This is in line with the goals of the Habitat Directive and Natura 2000 Network, which recommend prioritising the protection of populations / areas with Pan-European significance (Hoffman et al. 2004). However, establishing such areas for pond bats requires developing comparable data, based on extensive inventories, which are far from complete in most of the countries that acceded to the EU between 2004 and 2007 (cf. Limpens 2001). Acknowledgements: We are very grateful to Phil Richardson for correcting the English. We are also grateful to everybody who made their unpublished data available to this review (their contributions are acknowledged in the Annex by their initials): Gerben Achterkamp (GA), Sławomir Chmielewski (SC), Jakub Paweł Cygan (JPC), Robert Dróżdż (RD), Joanna Furmankiewicz (JF), Maciej Fuszara (MF), Judyta Gulatowska (JG), Mariusz Gwardjan (MG), Janusz Hejduk (JH), Maurycy Ignaczak (MI), Radosław Jaros (RJ), Anne Jifke Haarsma (AJH), Hans Huitema (HH), Krzysztof Kasprzyk (KK), Paweł Kmiecik (PK), Marek Kowalski (MK), Bart Kranstauber (BK), Ireneusz Krasnodębski (IK), Grzegorz Lesiński (GL), Peter H. C. Lina (PHCL), Czesław Nitecki (CN), Tomasz Nitkiewicz (TN), Agnieszka Ostrach-Kowalska (AO-K), Wojciech Pawenta (WP), Michał Piskorski (MP), Tomasz Postawa (TP), Agnieszka Przesmycka (AP), Ireneusz Ruczyński (IR), Aleksandra Szarlik (AS), Rafał Szkudlarek (RS), Marcin Urban (MU), Ben Verboom (BV), Michał Wojciechowski (MW), Zbigniew Wojciechowski (ZW), Agnieszka Wower (AW) and Sławomir Zalasa (SZ). In addition we would like to thank Joanna Furmankiewicz (Wrocław University) for her invaluable help during the radio-tracking study at the Jeleniewo nursery roost. References Alexandrowicz, S.W., A. Nadachowski, J. Rydlewski, P. Valde-Nowak & B.W. Wołoszyn Subfossil fauna from a cave in the Sobczański Gully (Pieniny Mts., Poland). Folia Quaternaria 56: Baagøe, H. J Danish bats (Mammalia: Chiroptera): Atlas and analysis of distribution, occurrence and abundance. Steenstrupia 26 (1): Bagrowska-Urbańczyk, E. & Z. Urbańczyk Structure and dynamics of a winter colony of bats. Acta Theriologica 28: Baranauskas, K New data on bats hibernating in underground sites in Vilnius, Lithuania. Acta Zoologica Lithuanica 16: Bernard, R. & J. Samoląg Dekady Spisu Nietoperzy w Strzalinach (północnozachodnia Polska). Nietoperze 3: Bolshakov, V.N. & O.L. Orlov The Species Composition of Bats Hibernating in Caves in the Central Ural Region. In: B.W. Wołoszyn (ed.). Approaches to Biogeography and Ecology of Bats. Proceedings of the VIIIth EBRS Vol. 1: CIC ISEZ PAN, Kraków, Poland. Chistyakov, D.V Present State of the Bat Community Hibernating in Artificial Caves near St. Petersburg. In: B.W. Wołoszyn (ed.). Distribution, Ecology, Paleontology and Systematics of Bats: Proceedings of the VIIIth EBRS. Vol. 2. CIC ISEZ PAN, Kraków, Poland. Ciechanowski / Lutra (2):

14 Ciechanowski, M. & A. Przesmycka Stwierdzenie nocka łydkowłosego Myotis dasycneme (Boie, 1825) i nocka wąsatka Myotis mystacinus (Kuhl, 1817) w Gdańsku. Nietoperze 2: Ciechanowski, M., A. Przesmycka & K. Sachanowicz 2006a. Species composition, spatial distribution and population dynamics of bats hibernating in Wisłoujście Fortress. Lynx 37: Ciechanowski, M., A. Przesmycka & K. Sachanowicz 2006b. Nietoperze (Chiroptera) Wdzydzkiego Parku Krajobrazowego. Parki Narodowe i Rezerwaty Przyrody 25: Ciechanowski, M., A. Przesmycka, A. Benedycka, A. Biała & K. Sachanowicz 2003a. Stwierdzenia rozrodu nocka łydkowłosego Myotis dasycneme (Boie, 1825) na Pojezierzu Pomorskim. Nietoperze 4: Ciechanowski, M., A. Zwolicki, M. Wojciechowski, A. Benedycka & A. Biała 2003b. Ssaki (Mammalia) projektowanego rezerwatu Torfowiska Wiszące nad Jeziorem Jaczno (Pojezierze Suwalskie) i jego otoczenia. Parki Narodowe i Rezerwaty Przyrody 22: Ciechanowski, M., L. Koziróg, J. Duriasz, A. Przesmycka, A. Świątkowska, J. Kisicka & K. Kasprzyk Bat fauna of the Iława Lakeland Landscape Park (northern Poland). Myotis 40: Čmak, J Ssaki (Mammalia) w biotopach Chełmowej Góry na tle fauny ssaków Świętokrzyskiego Parku Narodowego. Folia Forestalia Polonica 14: Dzięgielewska, M Zimowe liczenia nietoperzy w Szczecinie w latach Nietoperze 3: Encarnação, J.A., U. Kierdorf, D. Holweg, U. Jasnoch & V. Wolters Sex-related differences in roost-site selection by Daubenton s bats Myotis daubentonii during the nursery period. Mammal Review 35: Furmankiewicz, J. & J. Górniak Seasonal changes in number and diversity of bat species (Chiroptera) in the Stolec mine (SW Poland). Przyroda Sudetów Zachodnich Suppl. 2: Furmankiewicz, J. & M. Furmankiewicz Bats hibernating in the natural caves in the Polish part of the Sudetes. Przyroda Sudetów Zachodnich Suppl. 2: Furmankiewicz, J., M. Furmankiewicz & S. Telatyński Nowe obserwacje nocka łydkowłosego Myotis dasycneme (Boie, 1825) w polskiej części Sudetów Zachodnich. Przyroda Sudetów Zachodnich 4: Fuszara, E., M. Fuszara & M. Wojciechowski Monitoring liczebności nietoperzy w zimowiskach na Pomorzu Mazurskim w latach Nietoperze 3: Gas, A. & T. Postawa Bat fauna of the Studnisko Cave. Studia Chiropterologica 2: Gawlak, A Stanowisko nocka łydkowłosego Myotis dasycneme (Boie, 1825) w Poznaniu. Przegląd Przyrodniczy 7 (1): Gawlak, A., G. Wojtaszyn & M. Gmaj Zimowe spisy nietoperzy na Pomorzu Środkowym. Nietoperze 3: Godawa, J. & B.W. Wołoszyn Nietoperze (Mammalia: Chiroptera) Ojcowskiego Parku Narodowego. Prądnik. Prace Muzeum Szafera 1: Godawa, J Zmiany w faunie nietoperzy Ojcowskiego Parku Narodowego w latach : seria badań kontrolnych w latach Prądnik. Prace Muzeum Szafera 9: Grzywiński, W., J. Nowak & A. Węgiel Nietoperze Ojcowskiego Parku Narodowego - stan poznania. In: J. Partyka (ed.). Zróżnicowanie i przemiany środowiska przyrodniczo-kulturowego Wyżyny Krakowsko-Częstochowskiej. Tom I. Przyroda: Ojcowski Park Narodowy, Ojców, Poland Gulatowska, J. & M. Kowalski Największe zimowisko nocka łydkowłosego Myotis dasycneme na Nizinie Mazowieckiej. Nietoperze 5: Haitlinger, R Nietoperze Dolnego Śląska. Przegląd Zoologiczny 20 (1): Harmata, W Wyniki obrączkowania nietoperzy w Polsce w latach In: B. W. Wołoszyn (ed.). Aktualne problemy ochrony nietoperzy w Polsce: CIC ISEZ PAN, Kraków, Poland. Hejduk, J. & G. Radzicki Dynamika liczebności nietoperzy zimujących w Jaskini Szachownica (w sezonach 1993/94 i 1994/95). In: B. W. Wołoszyn (ed.). Aktualne problemy ochrony nietoperzy w Polsce: CIC ISEZ PAN, Kraków, Poland. Hoffman, J., A. Nowakowski & D. Metera 2004 (ed.). Integrating Natura Rural Development and Agri-Environmental Programmes in Central Europe. IUCN Foundation Poland, Warsaw, Poland. Horáček, I. & V. Hanák Distributional status of Myotis dasycneme. In: I. Horáček, V. Hanák & J. Gaisler (eds.). European Bat Research 1987: Charles University Press, Praha, Czech Republic. Horáček, I Myotis dasycneme (Boie, 1825). In: A.J. Mitchell-Jones, G. Amori, W. Bogdanowicz, B. Kryštufek, P.J.H. Reinjders, F. Spitzenberger, M. Stubbe, J.B.M. Thissen, V. Vohralik & J. Zima (eds.). The Atlas of European Mammals: Academic Press, London, UK. 120 Ciechanowski / Lutra (2):

15 Hutson, A.M Migrants, vagrants and stowaways. Bat News 77: 4-5. Hutson, A.M., S.P. Mickleburgh & P.A. Racey Microchiropteran Bats. Global Status Survey and Conservation Action Plan. IUCN/SSC Chiroptera Specialist Group, Gland, Switzerland and Cambridge, UK. Ignaczak, M. & J. Manias Nietoperze zasiedlające mosty w środkowej Polsce. Nietoperze 5: Jachno, J Dalszy ciąg zapisków fauniczych w Sandomierskiej Puszczy. Sprawozdanie Komisji Fizjograficznej 2: Janyszak, S. & M. Jurczyszyn Nocek łydkowłosy na Pomorzu Zachodnim. Biuletyn CIC 2 (4): 5. Jurczyszyn, M Nietoperze (Chiroptera) Roztoczańskiego Parku Narodowego i otuliny. Fragmenta Faunistica 37: Kasprzyk, K. & I. Ruczyński The structure of bat communities roosting in bird nest boxes in two pine monocultures in Poland. Folia Zoologica 50: Kliś, T., J. Furmankiewicz & T. Kokurewicz Zmiany liczebności i składu gatunkowego nietoperzy hibernujących w jaskiniach góry Połom (Góry Kaczawskie, Sudety Zachodnie) w latach Studia Chiropterologica 2: Kowalski, K Materiały do rozmieszczenia i ekologii nietoperzy jaskiniowych w Polsce. Fragmenta Faunistica 6: Kowalski, K Nasze nietoperze i ich ochrona. Polska Akademia Nauk, Zakład Ochrony Przyrody, Kraków, Poland. Kowalski, K Insectivores, bats and rodents from Early Pleistocene bone breccia of Podlesice near Kroczyce (Poland). Acta Palaeontologica Polonica 1: Kowalski, K., A. Krzanowski & R.J. Wojtusiak Report on bat-banding in Poland in the years Acta Theriologica 1: Kowalski, K., M. Młynarski, A. Wiktor & B.W. Wołoszyn Postglacjalna fauna z Józefowa, pow. Biłgoraj. Folia Quaternaria 14: Kowalski, M. & G. Lesiński Fauna drobnych ssaków w Janowie (woj. stołeczne) na podstawie analizy zrzutek płomykówki (Tyto alba Scop.). Przegląd Zoologiczny 30: Kowalski, M. & G. Lesiński The food of the tawny owl (Strix aluco L.) from near a bat cave in Poland. Bonner Zoologische Beiträge 41: Kowalski, M. & G. Lesiński 1994a. Bats occupying nest boxes for birds and bats in Poland. Nyctalus (N.F.) 5: Kowalski, M. & G. Lesiński Skład gatunkowy i wybiórczość kryjówek nietoperzy w Puszczy Kampinoskiej. Przegląd Przyrodniczy 6 (2): Kowalski, M. & G. Lesiński Nietoperze w diecie sów na Nizinie Mazowieckiej i Podlaskiej. Nietoperze 3: Kowalski, M. & R. Dróżdż Zimowy monitoring nietoperzy w sztucznej jaskini w Bochotnicy w latach Nietoperze 3 (1): Kowalski, M., G. Lesiński & M. Ignaczak Zimowy monitoring nietoperzy w jaskiniach na Wyżynie Wieluńskiej w latach Nietoperze 3 (1): Kowalski, M., G. Lesiński, B. Wojtowicz & T. Nitkiewicz Zimowe stanowiska nocka łydkowłosego Myotis dasycneme (Boie, 1825) w północnej części Podlasia. Nietoperze 4 (2): Kowalski, M., I. Krasnodębski, K. Sachanowicz, R. Dróżdż & B. Wojtowicz Skład gatunkowy, wybiórczość kryjówek i miejsc żerowania nietoperzy w Puszczy Kozienickiej. Kulon 1 (1-2): Kozakiewicz, K. & M. Strzałka Dynamika liczebności nietoperzy hibernujących w Jaskini Nietoperzowej w sezonie zimowym 1994/95. In: B.W. Wołoszyn (ed.). Aktualne problemy ochrony nietoperzy w Polsce: CIC ISEZ PAN, Kraków, Poland. Kozikowski, A. & L. Niedzielski Nietoperze (Chiroptera) wprzęgnięte do walki ze szkodliwymi owadami leśnymi. Sylwan 48 (1): Krzanowski, A Nietoperze (Chiroptera) Puław. Wykaz gatunków i uwagi biologiczne. Acta Theriologica 1 (4): Krzanowski, A. 1959a. Some major aspect of population turnover in wintering bats in the cave in Puławy (Poland). Acta Theriologica 3: Krzanowski, A. 1959b. Unusual summer hiding-place of bats. Acta Theriologica 2: Krzanowski, A Investigations of flights of Polish bats, mainly Myotis myotis (Borkhausen 1797). Acta Theriologica 4: Krzanowski, A Kompletna lista nietoperzy Puław. Przegląd Zoologiczny 7: Lesiński, G. & M. Kowalski Zimowy monitoring nietoperzy w Dolinie Narwi i Biebrzy w latach Nietoperze 3: Lesiński, G Nietoperze jaskiń Wyżyny Wieluńskiej. Przegląd Zoologiczny 27 (4): Lesiński, G Skład gatunkowy i liczebność nietoperzy w fortach modlińskich w ciągu roku. Przegląd Zoologiczny 32 (4): Ciechanowski / Lutra (2):

16 Lesiński, G Spis nietoperzy zimujących w Nietoperku z grudnia Wszechświat 89 (9): 209. Lesiński, G Nocek łydkowłosy w ptasich budkach. Wszechświat 90: 183. Lesiński, G Nietoperze Kotliny Biebrzańskiej i terenów przyległych. Parki Narodowe i Rezerwaty Przyrody 20 (2): Lesiński, G Nietoperze. In: Andrzejewski R. (ed.). Kampinoski Park Narodowy Vol. I: Kampinoski Park Narodowy, Izabelin, Poland. Lesiński, G Bat road casualties and factors determining their number. Mammalia 71: Lesiński, G., M. Kowalski, J. Domański, R. Dzięciołowski, K. Laskowska-Dzięciołowska & M. Dzięgielewska The importance of small cellars to bat hibernation in Poland. Mammalia 68: Limpens, H.J.G.A Assesing the European distribution of the pond bat (Myotis dasycneme) using bat detectors and other survey methods. Nietoperze 2: Limpens, H.J.G.A., P.H.C. Lina & A.M. Hutson Action plan for the conservation of the pond bat (Myotis dasycneme) in Europe. Council of Europe Publishing, Strasbourg, France. Lutsar, L., M. Masing & L. Poots Changes in the number of hibernating bats in the caves of Piusa (Estonia), Folia Theriologica Estonica 5: Łupicki, D. & M. Kowalcze-Łupicka Dominacja gatunków nietoperzy odławianych przy głównym wjeździe do podziemi MRU przed i po zmianach konstrukcji kraty zamykającej w 1995 r. In: R. Dzięciołowski (ed.). Materiały Konferencyjne, XIII Ogólnopolska Konferencja Chiropterologiczna, Błażejewko, 5-7 XI 1999: 31. PTOP Salamandra, Poznań, Poland. Łupicki, D., R. Szkudlarek, P. Schick & I. Dudek Wykorzystanie obiektów podziemnych przez nietoperze w rezerwacie Nietoperek w okresie jesiennym. Nietoperze 2 (1): Madeyska, T Środowisko człowieka w środkowym i górnym paleolicie na ziemiach polskich w świetle badań geologicznych. Studia Geologica Polonica 69: Mickevičiene, I., E. Mickevičius & K. Baranauskas 1999: Notes on summer distribution of bats in Lithuania. Acta Zoologica Lithuanica 9: Mikusek, R. & B. Pikulska Ssaki Parku Narodowego Gór Stołowych. Szczeliniec 3: Mleczek, T Zimowe spisy nietoperzy na Pogórzu Karpackim w latach Nietoperze 3: Mysłajek, R.W Nietoperze Chiroptera Parku Krajobrazowego Beskidu Małego. Nietoperze 3: Mysłajek, R.W., K. Henel & S. Nowak Nietoperze Chiroptera rezerwatu Łężczok koło Raciborza. Nietoperze 3: Nikodem, Z Materiały do fauny nietoperzy (Chiroptera) Lubelszczyzny. Przegląd Zoologiczny 26 (2): Nowak, J. & K. Kozakiewicz Zimowe spisy nietoperzy na Wyżynie Krakowskiej w latach Studia Chiropterologica 1: Nowak, J., A. Gawlak & G. Wojtaszyn Nocek łydkowłosy Myotis dasycneme (Boie, 1825) w Tatrach. Nietoperze 2 (1): Nowak, J., W. Grzywiński & M. Wieczorek Zimowe spisy nietoperzy na Wyżynie Krakowskiej w latach na tle 15 lat badań. Prądnik. Prace Muzeum Szafera 13: Ochman, K. & B.W. Wołoszyn Bats (Chiroptera). In: P. Valde-Nowak, A. Nadachowski & T. Madeyska (eds.). Obłazowa Cave. Human activity, stratigraphy and palaeoenvironment: Institute of Archaeology and Ethnology, Polish Academy of Sciences, Kraków, Poland. Pauža, D. H. & N. Paužiene Distribution and Status of Lithuanian Bats. Proceedings of Latvian Academy of Sciences, Section B 52 (1/2): Pax, F Säugetiere. In: Wirbeltierfauna von Schlesien: Gebrüder Borntraeger. Berlin, Germany. Pētersons, G. & V. Vintulis Distribution and Status of Bats in Latvia. Proceedings of Latvian Academy of Sciences, Section B 52 (1/2): Piksa, K. & B.W. Wołoszyn The postglacial bat remains from the Polish Tatra caves. Lynx (n.s.) 32: Piksa, K. & J. Nowak The Bat Fauna of the Polish Tatra Caves. In: B.W. Wołoszyn (ed.). Approaches to Biogeography and Ecology of Bats: Proceedings of the VIIIth EBRS. Vol. 1. CIC ISEZ PAN, Kraków, Poland. Pjenčák, P., S. Danko & S. Matis Netopiere Tatranského národného parku a širšieho okolia. Vespertilio 7: Postawa, T. & A. Gas Fauna nietoperzy Wigierskiego Parku Narodowego (północnowschodnia Polska). Studia Chiropterologica 3-4: Postawa, T. & J. Zygmunt Zmiany liczebności nietoperzy (Chiroptera) w Jaskiniach Wyżyny Częstochowskiej w latach Studia Chiropterologica 1: Ciechanowski / Lutra (2):

17 Postawa, T Changes in bat fauna during the Middle and Late Holocene as exemplified by thanatocoenoses dated with 14 C AMS from Kraków-Częstochowa Upland caves, Poland. Acta Chiropterologica 6: Rachwald, A., P. Boratyński & W. K. Nowakowski Species composition and night-time activity of bats flying over rivers in Białowieża Primeval Forest (Eastern Poland). Acta Theriologica 46: Reinhold, J.O., A.J. Hendriks, L.K. Slager & M. Ohm Transfer of microcontaminants from sediment to chironomids and the risk for the Pond bat Myotis dasycneme (Chiroptera) preying on them. Aquatic Ecology 33: Ruprecht, A.L Bats (Chiroptera) as constituents of the food of barn owls Tyto alba in Poland. Ibis 121: Ruprecht, A.L Food of the Barn owl, Tyto alba guttata (C.L.Bh.) from Kujawy. Acta Ornithologica 19: Ruprecht, A.L Myotis dasycneme (Boie, 1825). In: Z. Pucek & J. Raczyński (ed.). Atlas of Polish Mammals: 69-70, map no. 43. Polish Scientific Publishers, Warszawa, Poland. Ruprecht, A.L Nietoperze (Chiroptera) w składzie pokarmu sów z Puszczy Nadnoteckiej. Przegląd Zoologiczny 34: Sachanowicz, K. & I. Krasnodębski Skład gatunkowy i antropogeniczne kryjówki nietoperzy w Lasach Łukowskich. Nietoperze 4: Sachanowicz, K. & M. Ciechanowski Nietoperze Polski. Multico Oficyna Wydawnicza, Warszawa, Poland. Sachanowicz, K Zimowe stanowiska nietoperzy Chiroptera w południowej części Wysoczyzny Drohiczyńskiej i w regionie Podlaskiego Przełomu Bugu (Nizina Podlaska). Nietoperze 4 (1): Sachanowicz, K Structure and dynamics of the bat assemblage inhabiting military bunkers. Nyctalus (N.F) 12 (1): Sachanowicz, K., M. Ciechanowski & K. Piksa Distribution patterns, species richness and status of bats in Poland. Vespertilio 9-10: Sitowski, L Przyczynek do znajomości fauny Parku Narodowego w Pieninach. Ochrona Przyrody 18: Stebbings, R.E. & F. Griffith Distribution and status of bats in Europe. Institute of Terrestrial Ecology, Abbots Ripton, Huntingdon, UK. Strelkov, P.P. & V.J. Iljin The bats (Chiroptera: Vespertilionidae) of the south Middle Volga and Lower Volga provinces. In: P.P. Strelkov (ed.). Fauna, systematic and evolution of mammals (Rodentia, Chiroptera): Proceedings of the Zoological Institute, Leningrad, Russia. Styka, R Fauna drobnych ssaków Sobiborskiego Parku Krajobrazowego na podstawie analizy zrzutek puszczyka Strix aluco i płomykówki Tyto alba. In: Materiały VIII Ogólnopolskiej Konferencji Teriologicznej Bioróżnorodność i ochrona ssaków : KUL & UMCS, Lublin, Poland. Szkudlarek, R. & M. Dzięgielewska Nietoperze w rezerwacie Bielinek wstępne wyniki badań. Folia Universitatis Agriculturae Stetinensis 231 (92): Szkudlarek, R. & R. Paszkiewicz Zimowe stanowiska rzadkich gatunków nietoperzy w Sudetach Zachodnich, Przyroda Sudetów Zachodnich 2: Szkudlarek, R., R. Paszkiewicz, T. Blohm, E. Nowak & D. Łupicki Bunkry Ziemi Lubuskiej jako schronienia nietoperzy. Nietoperze 2 (1): Urbańczyk, Z. & Z. Gólski Zimowe spisy nietoperzy na Ziemi Lubuskiej w latach In: B.W. Wołoszyn (ed.). Zimowe spisy nietoperzy w Polsce: Wyniki i ocena skuteczności: CIC ISEZ PAN, Kraków, Poland. Van de Sijpe, M., B. Vandendriessche, P. Voet, J. Vandenberghe, J. Duyck, E. Naeyaert, M. Manhaeve & E. Martens Summer distribution of the Pond bat Myotis dasycneme (Chiroptera, Vespertilionidae) in the west of Flanders (Belgium) with regard to water quality. Mammalia 68: Verboom, B Winterverblijven voor vleermuizen in Limburg. VZZ report Zoogdiervereniging VZZ, Arnhem, The Netherlands. Wałecki, A Fauna zwierząt ssących Warszawy i jej stosunek do fauny całego kraju. Pamiętnik Fizyograficzny 1 (3): Węgiel, A., R. Szkudlarek & T. Gottfried Skład gatunkowy, aktywność i struktura populacji nietoperzy odławianych latem przy otworach jaskiń w Beskidach. Nietoperze 5 (1-2): Wojciechowski, M., K. Kasprzyk & M. Jefimow Pierwsze stwierdzenie kolonii rozrodczej nocka łydkowłosego Myotis dasycneme (Boie, 1925) na terenie Polski. In: R. Dzięciołowski (ed.). Materiały Konferencyjne, XIII Ogólnopolska Konferencja Chiropterologiczna, Błażejewko, 5-7 XI 1999: 46. PTOP Salamandra. Poznań, Poland. Wojtaszyn, G Nietoperze Pojezierza Wałeckiego. Przegląd Przyrodniczy 13 (1-2): Ciechanowski / Lutra (2):

18 Wojtaszyn, G., T. Rutkowski & W. Stephan Nowe stanowisko nocka łydkowłosego Myotis dasycneme (Boie, 1825) na Pobrzeżu Bałtyku. Chrońmy Przyrodę Ojczystą 62 (2): Wołoszyn, B.W Nietoperze z jaskiń Gór Świętokrzyskich. Przegląd Zoologiczny 6 (2): Wołoszyn, B.W Nowe obserwacje nad nietoperzami z jaskiń Gór Świętokrzyskich. Przegląd Zoologiczny 8 (3): Wołoszyn, B.W Badania nietoperzy Dolnego Śląska, Przegląd Zoologiczny 12 (2): Wołoszyn, B.W Pliocene and pleistocene bats of Poland. Acta Palaeontologica Polonica 32: Wołoszyn, B.W Nietoperze Chiroptera. In: K. Kowalski (ed.). Historia i ewolucja lądowej fauny Polski. Folia Quaternaria 59-60: Wołoszyn, B.W Dekady Spisu Nietoperzy w Polsce z perspektywy 5-ciu lat badań ( ) Podsumowanie. In: B.W.Wołoszyn (ed.). Zimowe spisy nietoperzy w Polsce: Wyniki i ocena skuteczności: CIC ISEZ PAN, Kraków, Poland. Wołoszyn, B.W Myotis dasycneme (Boie, 1825). Nocek łydkowłosy. In: Z. Głowaciński (ed.). Polska czerwona księga zwierząt. Kręgowce: Państwowe Wydawnictwo Rolnicze i Leśne, Warszawa, Poland. Żmudziński, L Rekolonizacja głębokiego dna Wdzydz Południowych dowodem poprawy czystości jeziora. Przegląd Przyrodniczy 8 (3): Samenvatting Zeldzaam of onderschat? - Het voorkomen van de meervleermuis (Myotis dasycneme) in Polen Dit artikel geeft een overzicht van het voorkomen van de meervleermuis (Myotis dasycneme) in Polen, gebaseerd op alle beschikbare gepubliceerde en ongepubliceerde gegevens. De soort is waargenomen in 105 UTM-hokken, verspreid over een groot deel van Polen. Daarnaast zijn fossiele resten, vooral uit het Holoceen, gevonden in 13 UTM-hokken. Voortplanting is vastgesteld in 10 UTM-hokken, uitsluitend in het noordelijk merengebied en in de riviervalleien van het centrale deel van Polen. Er zijn slechts twee kraamkolonies gevonden, met respectievelijk 481 en 391 individuen. De overige zomerwaarnemingen (41 UTM-hokken) zijn verspreid door het land gedaan, ook in delen ver verwijderd van de voortplantingsgebieden. Het gaat hier om zichtwaarnemingen en vangsten van volwassen mannetjes, vrouwtjes die niet aan de voortplanting deelnamen of dieren waarvan het geslacht onbekend was. Zomerverblijven (n=21) waren vooral gevestigd in gebouwen, holtes in bruggen en vogel- en vleermuiskasten. Winterverblijfplaatsen (n=67) van de meervleermuis bevonden zich hoofdzakelijk in oude forten, kelders en grotten. Hiervan werd 63% (n=42) bezet door één dier, terwijl in slechts drie verblijven tien of meer (maximaal 34) dieren werden gevonden. In twee verblijven waar regelmatig werd geinventariseerd fluctueerden de aantallen meervleermuizen in de laatste jaren, in een ander winterverblijf namen de aantallen zelfs toe. Het merendeel van de beschikbare data (wintertellingen, vogel- en vleermuiskasten, braakballen van uilen, verkeersslachtoffers, fossiele resten, mistnet-vangsten) laat zien dat meervleermuizen minder dan 1% uitmaken van de vleermuizen in Polen. In enkele delen van het merengebied van noordelijk Polen is het percentage meervleermuizen in mistnet-vangsten echter hoger, 2,1-2,7%, in één geval zelfs 17,5%. Veel gebieden die in potentie geschikt zijn als leefgebied voor de meervleermuis zijn nog niet onderzocht. Dit suggereert dat de meervleermuis in Polen wel eens veel algemener zou kunnen zijn dan tot dusver werd gedacht. Received: 17 January 2007 Accepted: 7 October Ciechanowski / Lutra (2):

19 Annex 1. Breeding records of the pond bat in Poland. Explanations: r. river, l. lake(s), rs survey of roost, nt netting, tr radiotracking of individuals from Jeleniewo nursery, ad. adult, juv. juvenile, lact. lactating, act. sexually active. If no symbol is given, the numbers refer to the unsexed individuals or that no data about sex and age are available. Capital italics (e.g. AP) represent the initials of unpublished data owners (explained in the acknowledgements) or of the authors (MC Mateusz Ciechanowski, KS Konrad Sachanowicz, TK Tomasz Kokurewicz). UTM square Locality Method Date Number and status of XV 88 Lubnia, forester s lodge rs (nursery) Ciechanowski et al. 2003a bats Source MC & KS MC & AP MC & AP MC MC XV 89 Loryniec, Wda r. nt ad. lact., 1 juv., 1 juv. Ciechanowski et al. 2003a XV 89 Schodno l., outflow of Wda nt ad. lact. Ciechanowski et al juv. Ciechanowski et al juv. Ciechanowski et al XV 89 Płocice, Wda r. nt juv. Ciechanowski et al XV 98 Borsk, Wda r. nt ad. (4 lact.) Ciechanowski et al CC 33 Mikołajewice, old branch of Warta r. nt ad. lact. MI DC 99 Łomianki, on road found dead juv. Lesiński 2003 DE 05 stream between Januszewskie and Czerwica l. nt juv. Ciechanowski et al EB 51 Trześnia, church loft rs (nursery) large number Jachno 1868 EB 51 Sokolniki tree hole rs with juv. Jachno 1868 FF 20 Jeleniewo, church loft rs (nursery) MW ( ad. + juv.) Wojciechowski et al ( ad. + juv.) TK & JF TK & JF FF 20 Szelment Wielki l. tr TK & JF FF 21 Szurpiły l. tr TK & JF 1 nt: 1 juv., 1 juv.; 2 nt: 2 ad. lact., 1 juv.; 3 nt: 2 ad. (1 lact.), 1 juv.; 4 nt: 1 ad. lact., 1 juv.; 5 nt: 2 ad. lact., 1 juv. Ciechanowski / Lutra (2):

20 Annex 2. Non-breeding summer records of the pond bat in Poland 1. Explanations: r. river, l. lake(s), f.d. forest district, dt foraging or commuting bat, observed with ultrasound detector, rs survey of roost, nt netting, ad. adult, juv. juvenile, act. sexually active. If no symbol is given, the numbers refer to the unsexed individuals or that no data about sex and age are available. Capital italics (e.g. MW) represent the initials of unpublished data owners (explained in the acknowledgements) or of the authors (MC Mateusz Ciechanowski, KS Konrad Sachanowicz, TK Tomasz Kokurewicz). UTM Locality Method Date Number and Source square status of bats VU 46 Bielinek, Old Odra r. dt 1999 Szkudlarek & Dzięgielewska 2003 VV 60 Lower Odra Valley, canals dt 2001 RS WR 88 Kudowa Zdrój dt RS in: Mikusek & Pikulska 1999 WU 30 Międzyrzecz Fortified Front ( Nietoperek bat reserve), nt TK et al. undergrounds WU 96 Trzcianka, narrow between Długie l. & fish ponds nt Wojtaszyn 2002 XS 09 Tarchalice, old branch of Odra r. dt 2003 JF XS 46 Odra r. near Wrocław dt 2001 RS XV 76 Struga Siedmiu Jezior, between Płęsno and Skrzynka l. nt ad. MK CB 90 Wielkanocna Cave rs (1 ad.) MI CC 33 Jeziorsko reservoir nt ad. JH CC 33 Warta II, Mazur ox-bow lake, crevice in a bridge rs Ignaczak & Manias 2004 CC 42 Nobela, Niniwka r., crevice in a bridge rs Ignaczak & Manias 2004 CC 42 Kolasa, Niniwka r., crevice in a bridge rs Ignaczak & Manias 2004 CC 50 Brody, Grabia r., crevice in a bridge rs Ignaczak & Manias 2004 CD 82 Gociąż l., bird box rs ad. Kasprzyk & Ruczyński 2001 CE 95 Fabianki, Liwa r. nt Ciechanowski et al CE 99 Lake Drużno nature reserve dt MC nt CN (photo), MC (det.) CF 25 Wejherowo, Reda r. nt ad. MC ad. MC CF 52 Gdańsk-Sobieszewo, bat box rs ad. MC DB 87 Chlewiska, church rs ad. SC & MK DC 16 Rydwan and Okręt fish ponds nt ZW DC 20 Golesze, building loft rs (colony) up to 10 SZ DC 30 Tomaszów Mazowiecki, Pilica r. nt ad. JH, WP & RJ DC 71 Wólka Magierowa, Drzewiczka r. nt ad. act. MK, GL & AO-K DD 60 Kromnów f.d., Kampinos Forest, bird box rs ad. Kowalski & Lesiński 1995 DD 80 Łomna, church loft rs ad. Kowalski & Lesiński ad. Kowalski & Lesiński 1995 DD 80 Kiełpin, on road found dead Lesiński 2003 DE 05 canal between Twaruczek and Płaskie l. nt ad. Ciechanowski et al DE 21 Kostkowo f.d., bat box rs ad. JG & AS EB 69 Puławy, loft of forester s lodge, rs (colony) , 1 Krzanowski 1956 EC 08 Warszawa-Marymont, Kaskada Park, hollow Carpinus rs , 1 Wałecki 1881 betulus tree EC 15 Czersk, church loft rs IK ED 79 Motyka, bat box rs ad. MK & TN ED 99 Wierciszewo, campsite near Biebrza r. dt GA & AJH EE 35 Mikołajskie l. dt BV & HH EE 46 canal between Łuknajno and Śniardwy l. dt BV & HH FB 30 Zwierzyniec, city park nt ad. Jurczyszyn 1994 FC 70 Żłobek, barn rs (colony) MP FD 94 Białowieża Forest, Hwoźna r., bridge nt ad. IR FE 01 Gugny, bird boxes rs (3 ) Lesiński ad. Lesiński 2001 FE 15 Dręstwo l. dt BV & HH in: Lesiński 2001 FE 27 Rospuda r., bridge nt ad. BK FE 37 bridge between Necko and Białe l. dt BK 126 Ciechanowski / Lutra (2):